The seeding of an expanse of surface waters in the equatorial Pacific Ocean with low concentrations of dissolved iron triggered a massive phytoplankton bloom which consumed large quantities of carbon dioxide and nitrate that these microscopic plants cannot fully utilize under natural conditions. These and other observations provide unequivocal support for the hypothesis that phytoplankton growth in this oceanic region is limited by iron bioavailability.
The standard dilution technique can provide unbiased estimates of phytoplankton growth and microzooplankton grazing rates only when certain restrictive assumptions are met. The most important of these assumptions -that grazing impact varies in direct proportion to the dilution of grazer population density -can be easily violated when clearance rate of individual grazers and/or growth response of the grazer population vary significantly with food concentration over the course of the incubation. We have developed a modified protocol which now allows the dilution technique to be applied unambiguously, even when its original assumptions may be violated. The new protocol uses flow-cytometry measured disappearance of fluorescently labeled tracer cells (FLB or FLA) as an internal measure of 'relative grazing activity' in each dilution treatment. Coefficients of phytoplankton growth and mortality rates are determined from Model 11 regression analyses of 'net growth' versus 'relative grazing', rather than the usual Model I regressions of 'net growth' versus 'dilution factor'. Tests of this hybrid experimental design in the central equatorial Pacific during an EQPAC cruise in August 1992 gave results essentially identical to the standard dilution interpretation.
During the IronEx II experiment in the eastern equatorial Pacific (May to June 1995), the response of the microplankton community to mesoscale iron fertilization was followed using a combination of marker-pigment, microscopical and flow cytometric techniques. Phytoplankton standing stock bloomed dramatically over a period of 6 d following 3 iron additions of 2 and 1 nM, respectively. Carbon biomass in the patch increased by a factor of 4, chlorophyll a by about a factor of 16 and diatoms by > 70-fold relative to contemporaneous levels in the ambient community. The bloom then plateaued sharply and remained at a more or less constant level for 4 d, despite the addition of more iron (1 nM) and physiological indices (low C:chl a ratio and elevated photochemical quantum efficiency) suggesting that the cells were healthy and growing rapidly. Relatively large pennate diatoms (Nitzschia spp., median length 20 to 24 µm) dominated the patch bloom, with smaller pennate species and centric diatoms declining in relative importance. Heterotrophic bacteria increased at a slow rate (0.08 d -1 ) for >10 d during the experiment, as did heterotrophic nanoflagellates. There were also indications of enhanced cell size, cellular pigment content and possibly growth rates of small phytoplankton. Nonetheless, little difference was observed between the ambient community and the peak patch bloom with respect to the size composition of auto-and heterotrophic populations <10 µm in cell size. The relative constancy of the smaller size fractions, the sharp curtailment of net growth of the bloom after 6 d, and > 3-fold increase in large heterotrophic dinoflagellates and ciliates suggest that protistan grazers may have played an active role in controlling the phytoplankton response to increased iron availability.KEY WORDS: Diatom · Bloom · Phytoplankton pigments · Carbon:chlorophyll · Size structure · Heterotrophic protists
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